Differences in growth between offspring of anadromous and freshwater brown trout Salmo trutta

In this study, individual growth of juvenile offspring of anadromous and freshwater resident brown trout Salmo trutta and crosses between the two from the River Imsa, Norway, was estimated. The juveniles were incubated until hatching at two temperatures (±S.D. ), either 4.4 ± 1.5°C or 7.1 ± 0.6°C. Growth rate was estimated for 22 days in August–September when the fish on average were c. 8 g in wet mass, and the estimates were standardized to 1 g fish dry mass. Offspring of anadromous S. trutta grew better at both 15 and 18°C than offspring of freshwater resident S. trutta or offspring of crosses between the two S. trutta types. This difference appears not to result from a maternal effect because anadromous S. trutta grew better than the hybrids with anadromous mothers. Instead, this appears to be an inherited difference between the anadromous and the freshwater resident fish lending support to the hypothesis that anadromous and freshwater resident S. trutta in this river differ in genetic expression. Egg incubation temperature of S. trutta appeared not to influence the later growth as reported earlier from the studies of Atlantic salmon Salmo salar.     

Spawning characteristics of the anadromous brown trout in a small Swedish stream

The spawning pattern of the anadromous brown trout Salmo trutta was studied in Själsöån, a small stream in Gotland, Sweden, during eight winters between 1992–1993 and 1999–2000. The total length ( L _T) at spawning was normally distributed (185–890 mm) for females and multimodal for males (300, 400 and 550 mm most frequent length classes). Spawning males were significantly younger (2+ to 4+ years) than females (3+ to 5+ years). The sex‐ratio at the beginning and at the end of the spawning season favoured males. The mean ±  s . d . number of spawners was 70 ± 16 individuals per year. Migration into and out of the stream occurred between November and June. The highest number of spawning fish was found in the stream at the end of November or at the beginning of December. Migration mainly occurred during high water flow and at night. The majority of the females entered the stream and spawned the same (29·3% of all the females) or the next night (32·8% of all the females) while males may have stayed for 2 to 3 weeks (21·3% of all the males) in the stream before spawning. Males usually remained much longer in the stream (mean ±  s . d . 45 ± 56 days) than females (16 ± 30 days). Females lost more mass in the stream (mean ±  s . d . 17·3 ± 8·6%) than males (7·7 ± 9·6%). For both sexes, mass loss was positively correlated with the time spent in the stream. Only 7·3% of the males and 5·7% of the females occurred in the stream for >1 year. Spawning took place only during the night.     

An intensive study of allozyme variation in freshwater resident and anadromous trout,Salmo trutta L., in western Ireland*

Ten polymorphic enzyme coding loci were assayed using starch gel electrophoresis in anadromous (sea trout) and freshwater-resident trout, Salmo trutta, from two neighbouring river systems draining into Clew Bay in western Ireland. Samples were collected from the Burrishoole and from the neighbouring Newport river system. In the Burrishoole system, two samples were from above waterfalls impassable to migratory trout, two consisted of anadromous smolts or post-smolts and the remainder were freshwater-resident trout from freely accessible areas. There were large differences in gene frequency between trout from above impassable barriers and all others sampled, and these two groups also differed substantially from each other. More minor differences were evident between some other samples, particularly between trout from the upper lake and streams entering it and a stream associated with the lower lake. Two anadromous populations therefore occur in the Burrishoole system. In contrast, no differences were evident in two sampling years between anadromous trout and cohabiting freshwater-resident trout from the upper part of the Burrishoole system. Thus, in the upper part of this system, anadromous trout do not appear to comprise a separate population from non-migratory trout and interbreeding may occur.     

Distribution, growth and movement of River Usk brown trout (Salmo trutta)

The River Usk catchment in South Wales supports mainly freshwater resident brown trout, with few anadromous fish. Electric fishing and netting revealed that age-class distribution differed between main river and tributary habitats, the latter environment acting as a nursery area for young fish. Few fry were found at main river sites. Age-class distribution also differed between tributary systems, and possible reasons are discussed. Trapping experiments indicated that trout move to main river habitat at 2+ yr. Lengths at age (back-calculated from scale reading) were similar for main river and tributary resident fish at 1 and 2 year, but main river fish were larger at 3 and 4 yr. This habitat shift and size difference is discussed with reference to current angling regulations.     

Allozyme diversity in brown trout (Salmo trutta) from Central Spain: Genetic consequences of restocking

1.  The brown trout ( Salmo trutta ) represents one of the main freshwater resources in Spain, but habitat alterations and overharvesting have contributed to the decline or disappearance of numerous natural populations. In addition, reinforcement programs of wild populations based on releases of hatchery reared fish of exogenous origin compromise the conservation of remnant native trout resources.
2.  We present allozymic data from Central Spain trout populations including stocked and unstocked populations. Although the levels of genetic variation observed were low and affected by hatchery releases (p = 18.23%, Ho= 3.39%), they were within the range observed in other European areas.
3.  The effective introduction of hatchery reared fish is genetically homogenising the populations in the studied area and disturbing the ancestral pattern of genetic variation that distinguishes the Tajo and Duero basins. Within the eight natural populations analysed, seven had alleles assigned to the foreign trout. The introgression in these populations, following the LDH-5 * 90 allele frequency, ranged between 2% and 29.4%, but those values are not in concordance with the respective stocking effort undertaken in each population. Moreover, the release of hatchery-reared fish does not solve the problems related to the reduced size of wild populations and their recruitment instability.     

Phylogeographical lineages in brown trout (Salmo trutta): investigating microgeographical differentiation between native populations from Northern Spain

SUMMARY 1. The large microgeographical differentiation revealed by allozyme studies in brown trout ( Salmo trutta) populations is one of the most striking features of this species. Additionally, allozymes showed great genetic differences between Atlantic and Mediterranean populations on a macrogeographical scale.
2. This study was carried out in order to assess whether the great differences observed between Atlantic and Mediterranean populations persisted where the two are geographically close (the 'microgeographical scale'). Sixteen populations of brown trout, S. trutta , were screened for genetic variation at 25 allozyme loci. The sampling sites, which occupied a relatively small geographical area, were distributed across Cantabrian (Atlantic) and Mediterranean drainages in Northern Spain.
3. The neighbour-joining tree, inferred from Nei's genetic distance, showed that brown trout populations clustered into two different groups. These groups corresponded to the Cantabrian and the Mediterranean groups of populations, although no clear geographical pattern emerged within each of the groups. This geographical pattern is basically caused by significant differences in the frequency distribution of the CK-A1 * locus, with a higher frequency of * 115 in Cantabrian samples (0.586 ± 0.091) while allele * 100 was more frequent in Mediterranean samples (0.931 ± 0.038). In addition, this study revealed alleles exclusive to the Mediterranean and Cantabrian populations, agreeing with previous findings.
4. Genetic differentiation between Cantabrian and Mediterranean regions (14.19%) was similar to that estimated in Spain at a larger scale (13%), showing that most of the differences between the regions can be observed even in a small geographical area.     

Effects of sea trout stocking on the population genetics of landlocked brown trout, Salmo trutta L., in the Conwy River system, North Wales, U.K.

The effect of the introduction of fry of anadromous sea trout, Salmo trutta L., on the genetic integrity of landlocked brown trout populations was evaluated. Samples were taken from six brown trout populations from streams above impassable waterfalls in the Conwy river system (North Wales, U.K.) in 1989 and 1990. Three of these streams had no known stocking history and three had been stocked with sea trout fry from the lower Conwy system over the last few years. Representatives of these sea trout were collected from two streams in the lower Conwy system and from a hatchery. Allele frequencies at 13 loci, six of which were polymorphic, were determined by starch gel electrophoresis.
The stocked populations were intermediate in their allele frequencies between unstocked brown trout and sea trout samples. A principal component analysis suggested significant numbers of hybrids in all of the stocked streams. This shows that some of the introduced sea trout did not migrate down the falls to the sea, but stayed in fresh water and hybridized with the local population. The significance of this finding for the conservation of the genetic resource of brown trout stocks is discussed.     

Interpopulation variation in reproductive traits of anadromous female brown trout, Salmo trutta L.

We studied reproductive traits in nine anadromous brown trout, Salmo trutta L., populations in seven Norwegian rivers. Within populations we found a positive significant correlation between fish length and fecundity in all populations, and between fish length and egg diameter in five populations. There were significant differences in these relationships between populations from different rivers, and between populations from different locations within rivers. When adjusted for variation in fish length, mean fecundity and mean egg diameter showed a negative significant correlation among populations. The ratio of gonadal weight to somatic weight (gonadosomatic index) varied significantly among populations but was not associated with variation in fish length. Comparatively few large eggs were found in brown trout populations co-existing with several other fish species.     

Thermoregulatory behavior of brown trout,Salmo trutta

Brown trout, Salmo trutta, were allowed to thermoregulate individually in an electronic shuttlebox. Pooled data for 6 fish showed a diel pattern of preferred temperature, with a diurnal minimum of 10.3°C, an early nocturnal maximum of 13.7°C, a less pronounced mid-scotophase minimum of 11.7°C, and a secondary dawn maximum of 12.8°C, in a somewhat crepuscular pattern. The 24-hour mean preferendum was 12.2°C.     

Migratory timing, marine survival and growth of anadromous brown trout Salmo trutta in the River Imsa, Norway

The aim of the paper was to study sea migration, growth and survival of brown trout Salmo trutta of the River Imsa, 1976–2005. The migratory S. trutta were individually tagged and fish leaving or entering the river were monitored daily in traps located near the river mouth. The mean annual duration of the sea sojourn was 6–9 months for first-time migrants moving to sea between January and June. It was 8–18 months for those migrating to sea between July and December. Veteran migrants stayed 12 months or less at sea and most returned to the river in August. Early ascending fish stayed the longest in fresh water because most returned to sea in April to May. The day number of 50% cumulative smolt descent correlated negatively with mean water temperature in February to March and the February North Atlantic Oscillation index (NAOI). Mean annual sea growth during the first 2 years after smolting was higher for S. trutta spending the winter at sea than those wintering in the River Imsa. First year's sea growth was lower for S. trutta descending in spring than autumn. For first-time migrants, it correlated negatively with the February NAOI of the smolt year. Sea survival was higher for spring than autumn descending first-time migratory S. trutta with a maximum in May (14·9%). Number of anadromous S. trutta returning to the river increased linearly with the size of the cohort moving to sea, with no evidence of density-dependent sea mortality. Sea survival of S. trutta smolts moving to sea between January and June correlated positively both with the annual number of Atlantic Salmo salar smolts, the specific growth rate at sea, and time of seaward migration in spring. This is the first study indicating how environmental factors at the time of seaward migration influence the sea survival of S. trutta .     

Juxtaposed microsatellite systems as diagnostic markers for admixture: an empirical evaluation with brown trout (Salmo trutta) as model organism

A juxtaposed microsatellite system (JMS) is composed of two microsatellite repeat arrays separated by a sequence of less than 200 bp and more than 20 bp. This paper presents the first empirical evaluation of JMSs for the study of genetic admixture induced by man, with brown trout (Salmo trutta) as model organism. Two distinct admixture situations were studied: native populations from streams of the Atlantic basin and of the Mediterranean basin, respectively, all stocked with domestic strains originating from the Atlantic basin. For these two situations, we first evaluated by simulation the ability of JMSs to differentiate between alien alleles and naturally shared homoplasious or ancestral alleles, and thus to behave as diagnostic markers for admixture. Simulations indicated that JMSs are expected to be reliable diagnostic markers in most divergent (i.e. Mediterranean) populations and nonreliable diagnostic markers in most closely related (i.e. Atlantic) populations. Three JMSs were genotyped in domestic strains as well as in nonstocked and stocked populations of brown trout sampled in different rivers of the Mediterranean and Atlantic basins. The observed distributions of JMS haplotypes were consistent with simulation predictions confirming that JMSs were reliable diagnostic markers only over a given proportion of the species range, i.e. in substantially divergent populations. JMSs also reinforced the diagnostic character of three microsatellite sites for the studied Mediterranean populations. This last result is consistent with our simulation results which showed that, although much less frequently than at JMSs, diagnostic markers are likely to be found at single site microsatellites provided that the native Mediterranean population has a sufficiently small effective population size. For each population of the Mediterranean basin admixture coefficients did not differ significantly across JMSs and mean admixture coefficients sometimes differ among populations. The interpretation of the origin of JMS haplotypes based on the allele length variants was supported by nucleotide sequence analysis.     

Genetic detection of sex-specific dispersal in historical and contemporary populations of anadromous brown trout Salmo trutta

The study of sex-biased dispersal has attracted considerable attention in birds and mammals, but less in other taxa, including fishes. We analysed sex-specific dispersal in historical (1910s and 1950s) and contemporary (1990s) samples of anadromous brown trout Salmo trutta. We tested the hypothesis that dispersal is unbiased using information from microsatellite DNA and applying an assignment index for 11 temporally and spatially separated samples. Our results are most consistent with brown trout dispersal being male biased, and provide no evidence of female bias. We found no evidence that dispersal patterns changed over time, indicating that stocking with hatchery strains did not affect sex-specific dispersal.     

The importance of being familiar: individual recognition and social behavior in sea trout (Salmo trutta)

This study addressed two hypotheses: (1) that individual recognitionis used to reduce the cost of contesting resources in sea troutand (2) that the hatchery environment selects against individualrecognition. Predictions from these hypotheses were tested inlaboratory experiments where behavior and growth were studiedin juvenile sea trout. In tact groups of familiar fish werecompared with groups that contained both familiar and unfamiliarindividuals. In general, the results were in agreement withthe first hypothesis. Familiar fish in intact groups had morestable dominance ranks, higher food intake, and better utilizationof food. Familiarity also reduced the distance to the nearestneighbor. Furthermore, initiators of conflicts were more likelyto win against familiar fish than against strangers. These resultssupport game-theory-based hypotheses explaining the dear-enemyphenomenon as an effect of familiarity A picture emerges inwhich familiarity stabilizes the hierarchical structure of agroup and governs behavioral modifications that will promotefeeding and growth, in turn leading to higher fitness. The secondhypothesis, which predicted a reduced effect of individual recognitionin sea-ranched trout, was not strongly supported because familiarityaffected sea-ranched and wild trout similarly in most respects.However, familiarity was not beneficial for growth in sea-ranchedtrout, whereas it increased growth rate in wild fish. In addition,sea-ranched trout tended to maintain larger distances to theirnearest neighbors than did wild trout.     

High level of population genetic structuring in lake-run brown trout, Salmo trutta, of the Inari Basin, northern Finland

Rivers draining into (Lake) Inarijärvi, northern Finland, sustain a number of lake‐run brown trout, Salmo trutta, populations but, as with most lake‐run S. trutta systems, the level of population genetic structuring among populations is unknown. To address this and to assist fish stock management in the region, the population genetic structure of S. trutta collected from 28 sampling sites in rivers flowing into Inarijärvi was studied using 13 microsatellite loci. Populations were clustered into three separate groups, largely corresponding to geographic regions, with between‐region F_ST values ranging from 0·11 to 0·16. The significant differentiation observed between most populations within each region also implies that individual populations should be recognized as separate management units and actions to improve, and subsequently maintain, conditions for natural spawning should be prioritized. The results of this study further indicate that the trout from each of these regions may have different biological characteristics, such as local‐lake feeding behaviour among the western populations and strong isolation among the northern stocks. As a consequence, further research is warranted to better understand the level of ecological uniqueness of lake‐run S. trutta populations.     

Failure of a stocking policy, of hatchery reared brown trout, Salmo trutta L., in Asturias, Spain, detected using LDH-5* as a genetic marker

A useful genetic marker exists through the apparent fixation of the LDH-5 * 100 allele in wild populations of brown trout in rivers from Asturias, Spain, contrasted with the near fixation of the LDH-5 * 90 allele in hatchery populations used to stock these rivers. In sampling locations where natural reproduction occurred, the * 100 allele was found exclusively in all areas having no record of hatchery stocking. The * 100 allele also predominated in three stocked areas having natural reproduction; in two of these areas a few individuals of the 0 + age class were homozygous for the * 90 allele. These data indicated that all catchable and reproductive fish originated from indigenous populations and thus the policy of hatchery supplementation was a failure in these areas.     

The post‐spawning movements of migratory brown trout Salmo trutta L.

The post spawning behaviour of sea trout Salmo trutta was studied over a 2 year period in the river and estuary of the River Fowey, south‐west England. Forty‐five sea trout kelts were trapped immediately after spawning in December and intraperitoneally tagged with miniature acoustic transmitters. The subsequent emigration into coastal waters was monitored using acoustic receivers deployed throughout the river catchment. The levels of gill Na⁺K⁺ATPase activity in sea trout kelts sampled at the same time as the tagged fish were within the range of 2·5 to 4·5 μmol Pi per mg protein per h indicating that the post‐spawning fish were not physiologically adapted to salt water. The tagged kelts were resident in fresh water between 4 and 70 days before entering the estuary. Sixty two per cent of the tagged kelts subsequently migrated successfully into coastal waters, with a higher success rate for male fish (75%) than females (58%). There was a significant size related difference in the run‐timing of the kelts with the larger fish moving more quickly into coastal waters after spawning than smaller fish. Seaward migration within fresh water was predominantly nocturnal and generally occurred in conjunction with increasing river discharge and rising water temperature. Migration through the estuary continued to be predominantly nocturnal and occurred during an ebbing tide. Residency within the estuary varied amongst individuals although it was invariably short, with most fish moving out into coastal waters within one to two tidal cycles. Five tagged kelts returned from the coastal zone and re‐entered fresh water during April and June. Marine residence time varied between 89 and 145 days (mean 118 days) and the minimum estimated marine survival was c. 18%. One of these sea trout was subsequently recaptured after successfully spawning in the vicinity where it had been previously tagged demonstrating a degree of spawning site fidelity.     

Maintenance of an endemic lineage of brown trout (Salmo trutta) within the Duero river basin

The Iberian Peninsula contains diverse populations of freshwater fish, with major river basins comprising differentiated biogeographic units. The Duero River flows through the North‐Western Iberian Peninsula and is one of the most important rivers within the Iberian glacial refuge. Brown trout (Salmo trutta) populate this whole basin, and studies using both allozyme and microsatellite loci have detected a geographically sorted distribution of genetic variation in this species. In this work, sequences of the mitochondrial control region obtained from 299 brown trout from the Duero River were compared with other Iberian and European datasets. Two differentiated haplotype groups were detected inside the Duero River basin. One of them was related to the Atlantic (AT) lineage that is present in Northern European populations, whereas the other comprised an unique group that was restricted to the inner region of the basin. The amount of divergence of this Duero group from the other brown trout populations studied is consistent with a new trout lineage (Duero, DU) that is endemic to this river basin and that diverged from other Atlantic populations during the Pleistocene. The distribution of the DU and AT quaternary lineages in the Duero River was consistent with the ichthyological pattern described in the basin that originated during the Miocene–Pliocene. Evidence of selective processes that favour the haplotypes of the DU lineage may explain this discrepancy.     

Genetic structure of brown trout, salmo trutta l., at the southern limit of the distribution range of the anadromous form

Genetic variation at 33 protein loci was investigated in 41 wild brown trout populations from four river basins in Galicia (northwest Spain) to analyse the amount and distribution of genetic diversity in a marginal area, located in the distribution limit of the anadromous form of this species. The genetic diversity detected within populations (H between 0 and 6%) lies within the range quoted for this species in previous reports. The Mino, the most southern river basin analysed, showed a significantly lower genetic diversity and the highest genetic differentiation among the river basins studied. The hierarchical gene diversity analysis showed high population differentiation in a restricted area (GST = 27%), mostly due to differences among populations within basins (GSC = 22%). The reduction of GST observed when the isolated samples were excluded from the analysis (GST = 17%) showed the importance of habitat fragmentation on the heterogeneity detected. Gene flow among populations was comparatively evaluated by three indirect methods, which in general revealed low figures of absolute number of migrants per generation, slightly higher than 1. The gene flow among basins reflected a positive relationship with geographical distance. This trend was confirmed by the significant correlation observed between geographical and genetic distances, including all population pairs, which suggests a component of isolation by distance in brown trout genetic structure. Nevertheless, the nonsignificant intrabasin correlation demonstrates the complexity of genetic relationships among populations in this species. The model of genetic structure in brown trout is discussed in the light of the results obtained.     

Mitochondrial DNA variation in River Usk brown trout, Salmo trutta

Mitochondrial DNA (mtDNA) polymorphism in the essentially non-anadromous River Usk brown trout Salmo trutta population was investigated by restriction analysis. Following mtDNA extraction and purification on caesium chloride density gradients, monomorphic restriction profiles were obtained with Hae III , Hind III , Sau 3AI and Xbal . However, the restriction endonucleases Ava II and HinfIl proved informative. The distribution of four composite genotypes found within the Usk system was heterogeneous, and a fifth genotype appeared exclusively in an outgroup sample from the adjacent River Wye drainage. The source of the observed genetic variation is discussed in relation to estimated divergence times for Usk mtDNA genotypes and the stocking history of the catchment.